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New Insights Into the Anatomy of Extinct Paucituberculatan Marsupials

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New Insights Into the Anatomy of Extinct Paucituberculatan Marsupials Swiss J Palaeontol DOI 10.1007/s13358-014-0063-9 An exceptionally well-preserved skeleton of Palaeothentes from the Early Miocene of Patagonia, Argentina: new insights into the anatomy of extinct paucituberculatan marsupials Analia M. Forasiepi • Marcelo R. Sa´nchez-Villagra • Thomas Schmelzle • Sandrine Ladeve`ze • Richard F. Kay Received: 8 September 2013 / Accepted: 13 February 2014 Ó Akademie der Naturwissenschaften Schweiz (SCNAT) 2014 Abstract During the Cenozoic paucituberculatans were an anterior semicircular canal (SC) projecting slightly much more diverse taxonomically and ecomorphologically dorsally from the dorsal-most point of the posterior SC, and than the three extant genera of shrew-like marsupials. lateral and posterior SCs projecting laterally to the same Among paucituberculatans, palaeothentids were abundant level. On the basis of postcranial anatomy, previous studies during the Early Miocene, although most of the fossil have demonstrated that P. lemoinei was an agile cursorial remains consist of isolated teeth or fragmentary jaws. We form, an inference supported by study of the new post- describe a new and exceptional partial skeleton of Palaeo- cranial elements. thentes lemoinei (Palaeothentidae), collected from the Santa Cruz Formation (Santacrucian age, Early Miocene) Keywords Marsupialia Á Metatheria Á Cenozoic Á in Patagonia. Whereas the skull of P. lemoinei has more South America Á Skull Á Inner ear Á Postcranium plesiomorphic traits in the face, palate, and cranial vault than that of living paucituberculatans, the dental mor- Abbreviations phology is more derived. The osseous inner ear was examined using micro-CT scanning, revealing a cochlea Institutional abbreviations with 1.9 turns, the presence of a ‘‘second crus commune’’, FMNH Field Museum of Natural History, Chicago, USA IEEUACH Universidad Austral de Chile, A. M. Forasiepi Instituto de Ecologı´a y Evolucio´n, Instituto Argentino de Nivologı´a, Glaciologı´a y Ciencias Valdivia, Chile Ambientales, CCT-Mendoza, CONICET, Av. Ruiz Leal s/no, MACN-A Museo Argentino de Ciencias 5500 Mendoza, Argentina Naturales ‘‘Bernardino Rivadavia’’, A. M. Forasiepi Á M. R. Sa´nchez-Villagra (&) Ameghino Collection, Buenos Aires, Palaeontological Institute and Museum, University of Zu¨rich, Argentina Karl Schmid-Strasse 4, 8006 Zu¨rich, Switzerland MNHN-Pal-Bol-V Museo Nacional de Historia Natural, e-mail: [email protected] Vertebrates, La Paz, Bolivia T. Schmelzle MPM-PV Museo Regional Provincial ‘‘Padre Fleckensteinstraße 7, 74206 Bad Wimpfen, Germany M. Jesu´s Molina’’, Rı´o Gallegos, Argentina S. Ladeve`ze PU Princeton University collections at Muse´um national d’Histoire naturelle, UMR 7207 CR2P CNRS/ MNHN/UPMC, 8, rue Buffon, CP38, 75231 Paris Cedex 05, Yale University, USA France Anatomical abbreviations R. F. Kay (&) I/i Upper and lower incisor Department of Evolutionary Anthropology, Duke University, C/c Upper and lower canine Durham, NC 27708-03083, USA P/p Upper and lower premolar e-mail: [email protected] A. M. Forasiepi et al. M/m Upper and lower molar Villagra 2003; Asher et al. 2004; Nilsson et al. 2004, 2010; ASC Anterior semicircular canal Beck et al. 2008; Horovitz et al. 2008, 2009; Meredith et al. LSC Lateral semicircular canal 2008). Other studies located paucituberculatans at the base PSC Posterior semicircular canal of Marsupialia (e.g., Szalay 1994; Baker et al. 2004; Meredith et al. 2009, 2011) or provided the two alternatives (Beck 2008). Consequently, the understanding of the Introduction anatomy of fossil paucituberculatans has the potential to provide new data to interpret the anatomy and the character Paucituberculata is a clade of shrew-like marsupials evolution of the Australian marsupials or early marsupials. endemic to South America. The group includes seven Unfortunately, the fossil record of paucituberculatans is extant small and insectivorous species arranged in three mainly confined to teeth. genera: Caenolestes, Lestoros, and Rhyncholestes (Wilson Paucituberculatans are typically classified into the Lin- and Reeder 1993; Ojala-Barbour et al. 2013). Fossil naean families Caenolestidae, which includes the living members of this group were much more diverse taxo- taxa, and the extinct Pichipilidae, Abderitidae and Palaeo- nomically and ecomorphologically (Marshall 1980; Bown thentidae, and stem taxa (Marshall 1980; Bown and Fleagle and Fleagle 1993; Abello 2007; Abello and Candela 2010) 1993; Abello 2007, 2013; Goin et al. 2009; Abello 2013; and are known since the Early Eocene (Itaboraian age) Fig. 1). Palaeothentidae occurred from the late Oligocene (Goin et al. 2009; Abello 2013; Fig. 1). to the middle Miocene (Deseadan to Laventan ages) in The alpha taxonomy and the phylogenetic relationships Colombia, Bolivia, southern Argentina, and Chile (Mar- of paucituberculatans has been thoroughly revised (Abello shall 1980; Bown and Fleagle 1993; Dumont and Bown 2007, 2013) and continues to be studied (Goin et al. 2010; 1997; Abello 2007, 2013). Palaeothentids were abundant Abello and Rubilar-Rogers 2012). The discovery of early during the late Early Miocene (Santacrucian age), with paucituberculatans and closer relatives (Goin et al. 2007, Palaeothentes being the most speciose genus, with five 2009; Forasiepi et al. 2013) significantly increases the species: P. aratae, P. intermedius, P. lemoinei, P. minutus, understanding of the character evolution within this clade. and P. pascuali. Despite an extensive fossil record, com- Several studies based on molecular, morphological, and plete skulls and skeletons are rare. Most of the fossil combined data agree that paucituberculatans are the sister remains consist of isolated teeth, fragmentary mandibles taxa of Australidelphia (e.g., Horovitz and Sa´nchez- and fragmentary maxillae (Abello 2007). To date, four Fig. 1 Phylogenetic relationships of several paucituberculatans based younger): Itab Itaboraian, Cas Casamayoran, SRF Santa Rosa Fauna, on Abello (2013), indicating Palaeothentinae (the monophyletic group Tin Tinguirirican; Des Deseadan, Col Colhuehuapian, P Pinturan to Palaeothentes lemoinei belongs) in bold upper case. South fauna; S–F–C Santacrucian, Friasian, and Colloncuran, Lav Laventan, American Land Mammal Ages (S) are as follows (from older to Huay Huayquerian, Mon Montermosan Skeleton of the marsupial Palaeothentes lemoinei skulls of palaeothentids have been described. One belongs easier comparison. For the description, the lateral semi- to the Santacrucian palaeothentid Palaeothentes minutus circular canal was oriented horizontally and at the centre of (MACN-A 8271), and was damaged subsequent to its the anatomical structures (Graf and Klam 2006). The turns illustration by Ameghino (1887, p. 6; see also Marshall of the cochlea were counted following the protocol of West 1980, fig 20). Two other partial skulls assignable to the (1985). acdestid Acdestis oweni are also from the Santacrucian: specimens PU 15225 and FMNH 13160, the former described by Sinclair (1906)asPalaeothentes intermedius Systematic palaeontology and referred to Acdestis spegazzinii by Abello (2007). The fourth skull is from rocks of Laventan age of Bolivia Marsupialia Illiger 1811 (MNHN-Pal-Bol-V-4000) and was described as Acdestis Paucituberculata Ameghino 1894 maddeni (Goin et al. 2003). Palaeothentidae Sinclair 1906 This contribution describes and examines a new speci- Palaeothentes Ameghino 1887 men, comprising the skull and postcranial skeleton, of the P. lemoinei Ameghino 1887 palaeothentid P. lemoinei (MPM-PV 3566), collected from Holotype. MACN-A 3, fragment of right dentary with m1–m4. Puesto Estancia La Costa of the Estancia La Costa Mem- ber, Santa Cruz Formation (Santacrucian age). The speci- Occurrence and age. Santa Cruz, Pinturas, and Rı´o Frias men was recovered in 2003 by RFK during a joint formations, Early–Middle Miocene (Santacrucian–Friasian expedition from Museo de la Plata (Argentina) and Duke South American Land Mammal Ages), Patagonia, Argen- University (USA). This new specimen adds insights into tina and Chile (Marshall 1980, 1990; Bown and Fleagle the knowledge of the anatomy of extinct paucitubercula- 1993; Abello 2007). tans and is the core of the present study, focusing on the Referred specimen. MPM-PV 3566, skull with dentition, middle ear anatomy. We apply the techniques of CT complete left and fragment of right dentary with dentition, technology to study the internal osseous inner ear anatomy right petrosal, and postcranium (16 vertebrae, ribs, right pelvic of P. lemoinei and to undertake comparisons with other girdle, right femur, proximal fibula, right astragalus, left Metatheria. humerus, left ulna, both radii, metapodials, and phalanxes). Remarks. The combination of the following features per- Materials and methods mits allocation of MPM-PV 3566 to P. lemoinei (for a taxonomic revision of the genus and revised diagnosis see The nomenclature for the descriptions follows veterinary Abello 2007): wide crown bases in M1–M2, larger in molar text books (e.g., Schaller 1992) and particular publications size than the Santacrucian species P. intermedius, P. based on metatherians. The nomenclature for the skull pascuali, and P. minutus and smaller than in P. aratae follows Wible (2003), for the petrosal Wible (1990, 2003), (Abello 2007; Tables 18a–d), less steep molar-size gradi- for the inner ear Schmelzle et al. (2007), and for the den- ent from m1 to m4, and more protruding protocone on M1. tition Abello (2007, 2013). Additionally, P. lemoinei has the plesiomorphic
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